Units and levels of selection

In a standard Darwinian explanation, natural selection takes place at the level of the individual organism, i.e. some organisms enjoy a survival or reproduction advantage over others, which results in evolutionary change. In principle however, natural selection could operate at other hierarchical levels too, above and below that of the organism, for example the level of genes, cells, groups, colonies or even whole species. This possibility gives rise to the ‘levels of selection’ question in evolutionary biology. Group and colony-level selection have been proposed, originally by Darwin, as a means by which altruism can evolve. (In biology, ‘altruism’ refers to behaviour which entails a fitness cost to the individual so behaving, but benefits others.) Though this idea is still alive today, many theorists regard kin selection as a superior explanation for the existence of altruism. Kin selection arises from the fact that relatives share genes, so if an organism behaves altruistically towards its relatives, there is a greater than random chance that the beneficiary of the altruistic action will itself be an altruist. Kin selection is closely bound up with the ‘gene’s eye view’ of evolution, which holds that genes, not organisms, are the true beneficiaries of the evolutionary process. The gene’s eye approach to evolution, though heuristically valuable, does not in itself resolve the levels of selection question, because selection processes that occur at many hierarchical levels can all be seen from a gene’s eye viewpoint. In recent years, the levels of selection discussion has been re-invigorated, and subtly transformed, by the important new work on the ‘major evolutionary transitions’. These transitions occur when a number of free-living biological units, originally capable of surviving and reproducing alone, become integrated into a larger whole, giving rise to a new biological unit at a higher level of organization. Evolutionary transitions are intimately bound up with the levels of selection issue, because during a transition the potential exists for selection to operate simultaneously at two different hierarchical levels.